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(Myrmeciza Exsul) and White-Bellied Antbird (M ORNITOLOGIA NEOTROPICAL 16: 397–404, 2005 © The Neotropical Ornithological Society NONBREEDING TERRITORIAL BEHAVIOR OF TWO CONGENERIC ANTBIRDS, CHESTNUT-BACKED ANTBIRD (MYRMECIZA EXSUL) AND WHITE-BELLIED ANTBIRD (M. LONGIPES) Bridget J. M. Stutchbury1, Bonnie E. Woolfenden1 , Bradley C. Fedy2, & Eugene S. Morton3 1Department of Biology, York University, Toronto, Ontario M3J 1P3, Canada. E-mail: [email protected] 2Centre for Applied Conservation Research, Forest Sciences Centre, University of British Columbia, Vancouver, British Columbia V6T 1Z4, Canada. 3Conservation and Research Center, Smithsonian Institution, Front Royal, Virginia 22630, USA. Resumen. – Comportamiento territorial no-reproductivo en dos hormigueros congenéricos, el Hormiguero Dorsicastaño (Myrmeciza exsul) y el Hormiguero Ventriblanco (M. longipes). – Utili- zamos radiotelemetría durante la época no-reproductiva (seca) para comparar sistemáticamente los movi- mientos a corto plazo, cantos y comportamiento territorial de dos hormigueros que tienen ecología similar pero ocupan diferentes hábitat. Ambas especies permanecen en parejas y son territoriales a lo largo del año, pero el Hormiguero Dorsicastaño (Myrmeciza exsul) es común en hábitat secundarios y de borde, mientras el Hormiguero Ventriblanco (M. longipes) es común en bosque maduro. Aunque el Hormiguero Ventriblanco realizó incursiones fuera del territorio durante la época seca, no detectamos ninguna salida en el Hormiguero Dorsicastaño. Los Hormigueros Ventriblancos cantaron más y cubrieron menos área de su territorio por hora (0,01 ha) comparados con los Hormigueros Dorsicastaños (0.16 ha). Las parejas de Hormigueros Ventriblancos forrajearon usualmente a más de 5 m uno del otro y los duetos fueron regula- res, aunque de baja frecuencia en la mayoría de las parejas. En las parejas de Hormigueros Dorsicastaños, los individuos forrajearon más separados, frecuentemente a más de 20 m de distancia. Las hembras de Hormigueros Dorsicastaños cantaban ocasionalmente y cuando lo hacían, esto era inmediatamente des- pués del canto de su pareja. Discutimos como estas diferencias en comportamiento territorial y cantos reflejan una mayor estabilidad del alimento, y por lo tanto una mayor estabilidad territorial de la especie de bosque, el Hormiguero Dorsicastaño. Abstract. – We used radiotelemetry during the nonbreeding (dry) season to systematically compare the short-term movements, singing and territorial behavior of two antbirds that have a similar ecology but occupy different habitats. Both species are paired and territorial year round, but the White-bellied Antbird (Myrmeciza longipes) is common in second growth and edge habitat, while the Chestnut-backed Antbird (M. exsul) is found in mature forest. Although White-bellied Antbirds made off-territory forays during the dry season, we did not detect any off-territory forays in Chestnut-backed Antbirds. White-bellied Antbirds sang more and covered less area of their territories per hour (0.01 ha) compared with Chestnut-backed Antbirds (0.16 ha). Mated White-bellied antbirds often foraged < 5 m from each other and duetting occurred regularly, though at low frequency for most pairs. In Chestnut-backed Antbirds, pairs foraged far- ther apart and were often separated by over 20 m. Female Chestnut-backed Antbirds only occasionally sang and, when they did, it was immediately after their mate’s song. We discuss whether these differences 397 STUTCHBURY ET AL. in territorial behavior and singing reflect greater food stability, and hence greater territorial stability, in the forest-dwelling Chestnut-backed Antbird. Accepted 27 April 2005. Key words: Territory defense, radiotelemetry, foraging, duetting, song, off-territory forays, Chestnut- backed Antbird, White-bellied Antbird, Myrmeciza exsul, Myrmeciza longipes. INTRODUCTION in general, would be more restricted in the Chestnut-backed Antbird which occupies In tropical passerines, year-round defense of mature forest. territories by pairs is a common territorial sys- We were also interested in determining if tem but relatively little is known about song song rates differed as a function of habitat. rates, off-territory forays, and territory Song rates in year-round territorial passerines defense in these birds (Stutchbury & Morton are typically low relative to seasonally territo- 2001). We compared the singing behavior and rial temperate zone passerines (reviewed in movements of two congeneric antbirds in Stutchbury & Morton 2001). Little is known Panama. The White-bellied Antbird (Myrme- about interspecific variation in singing behav- ciza longipes) is common in second growth and ior among tropical passerines, because so few edge habitat (Wetmore 1972) while the Chest- studies have systematically documented song nut-backed Antbird (M. exsul) occupies forest rates. If song rate is driven largely by food habitat (Skutch 1969). Their song structures availability (e.g., Reid 1987, Cuthill & Mac- show acoustic adaptation to their different donald 1990), we would expect to find that habitats (Morton 1975). A recent molecular during the dry season Chestnut-backed Ant- phylogeny suggests that the genus Myrmeciza birds (forest habitat) have a higher song rate is paraphyletic (Irestedt et al. 2004); neverthe- than White-bellied Antbirds (drier second less, the species’ ecology is superficially simi- growth habitat). lar (other than habitat) since they defend territories year-round as pairs, forage on or METHODS near the ground, and are insectivorous but do not regularly follow army ant swarms or join White-bellied Antbirds were studied during mixed-species flocks. the dry season from January to April of 2000 White-bellied Antbirds, a second growth and 2001 at two sites in Parque Nacional de species, have low song rates and unstable ter- Soberania, Panama (09o07’N, 79o40’W), one ritories during the nonbreeding dry season in the town of Gamboa and the other site at (Fedy & Stutchbury 2004). White-bellied Ant- the entrance to the Pipeline Road (Fedy & birds make off-territory forays and some indi- Stutchbury 2004). In these two study sites, the viduals abandoned their territory temporarily average territory size was 0.7 ha and 2.3 ha, for several weeks or months, possibly as a respectively. Most birds were individually response to food shortage (Fedy & Stutch- color banded, and territory boundaries were bury 2004). Arthropod abundance in edge determined by mapping bird locations while and second growth habitat tends to exhibit they sang, noting locations of counter-singing greater seasonality than in moister habitats at borders, and by assessing responses to play- like mangrove and forest (Levings & Windsor back of recorded songs. Three types of vocal- 1982, Parrish & Sherry 1994, Poulin & Lefeb- izations were distinguished: songs, “chips” vre 1997). Therefore we expected to find that and “cheers” (Fedy & Stutchbury 2004). off-territory forays, and territorial movements In 2001, eight different pairs of White- 398 ANTBIRD TERRITORIALITY bellied Antbirds (and two other males) were male was radio-tagged were recorded when- fitted with radio-transmitters early in the dry ever the female vocalized or was seen by the season (January) and followed for the next 6– observer. 8 weeks. BD-2G transmitters (Holohil Sys- For analysis of vocalizations, we treated tems Ltd., Carp, Ontario) weighing about 2 g each radio-tracking session as an independent each were attached using a figure-eight har- unit, rather than each individual territory, ness (Rappole & Tipton 1991) made of light- because song rates were highly variable weight cotton embroidery floss. We between tracking sessions. For analysis of dis- determined a bird’s location using hand-held tances between pair members, we treated the Yagi antennas and maintained a distance of at pair distance at 15 min intervals during track- least 15 m from the birds. Singing data were ing sessions as independent points. For collected continuously during 1-h observation White-bellied Antbirds, where we had the periods. The distance separating members of most observations, we calculated the percent- a pair was recorded every 15 min and each age of observations at critical distances time when the pair sang a duet. We tracked (< 5 m, > 20 m) for each pair. each individual for an average of 11.4 h (SD = The rainfall patterns during the dry season 4.1, range 4–17) and obtained a total of 205 h were similar for 2001–2003 on Barro Colo- of radio-tracking data on individual White- rado Island, where meteorological records are bellied Antbirds. A subset of these data were available (http://striweb.si.edu/esp). Rainfall used for examining the movements of pairs. in February was 14.5, 10.2 and 15.7 mm, We had 51 h of radiotracking data where both respectively, and was 25.6, 46.6, and 2.8 mm members of a pair were radio-tagged (average in March. The length of the dry season was 6.1 h per pair, 4–9 h per pair). 136, 123, and 134 d in the three years of the Chestnut-backed Antbirds were studied in study, all of which are close to the 50 year February 2003 on Barro Colorado Island, average of 135 days. Panama (09o10’N, 79o50’W), which is approxi- mately 10 km from Gamboa. We color RESULTS banded eight males and two females on eight adjacent focal territories. To determine terri- Movements. In White-bellied Antbirds, 6/10 tory boundaries, we surveyed the trails in an males and 2/8 females made at least one off- approximately 10-ha area, in the vicinity of territory foray (as reported earlier, Fedy & the laboratory buildings, by listening for sing- Stutchbury 2004), and 4 of the 18 White-bel- ing birds and conducting regular song play- lied Antbirds tracked spent > 25% of their backs. Three types of vocalizations were time off-territory. Individuals did not sing distinguished: songs, “snarls” which are often while off-territory and visited from 1-3 neigh- used by the mated pair in answering each boring territories. In Chestnut-backed Ant- other, and “cli-dicks” which are typically used birds, none of the four males, or the female, as alarm notes (Skutch 1969). ever left their territory during a total of 24 h We radio-tagged five birds on four differ- of tracking.
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